Distribution of carbohydrate during grainfill in Leafy and

normal maize hybrids

C. J. Andrews1, L. M. Dwyer1, D. W. Stewart1, J.-A. Dugas2, and P. Bonn1
1EasternCereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Central Experimental Farm,
Ottawa Ontario K1A 0C6; and 2Glenn Seed Ltd., Blenheim Ontario N0P 1A0, Canada. Received 12 February
1999, accepted 29 September 1999.

Andrews, C. J., Dwyer, L. M., Stewart, D. W., Dugas, J.-A. and Bonn, P. 2000. Distribution of carbohydrate during grainfill
in Leafy and normal maize hybrids. Can. J. Plant Sci. 80: 87–95. The Leafy (Lfy) genotype in corn has extra leaves above the
ear in comparison to normal non-leafy genotypes and has been shown to increase yields in some geographical areas. In this study,
carbohydrate distribution in three Lfy hybrids (two of which were “staygreen”) and a normal check (Pioneer 3790) were analyzed
in a short-season area (2800 Crop Heat Units, suitable for 85 day Relative Maturity hybrids). Total sugar and starch were deter-
mined at all aboveground levels of leaf and stem on 1 August 1990 and this distribution was used to calculate total canopy carbo-
hydrates in two subsequent years from samplings of leaf and stem at only three or four canopy levels bracketing the ear level.
Husk, cob and kernel components were also analyzed at three sampling times during grainfill. Major differences in the 2 yr were
observed, but in general there was approximately twice the carbohydrate in the canopy at and above the ear in the Lfy genotypes
compared to the check. There was also more carbohydrate in the husk and cob in the Lfy lines, but substantially less carbohydrate
below the ear in these lines. Rates of grainfill were generally higher in the Lfy hybrids, but the rate of Lfy hybrid A, the early
senescent hybrid, was slower in the cooler growing season. Despite the greater amount of carbohydrate in the Lfy hybrids in com-
parison to the traditional check, their grain yields were not greatly increased, indicating that the kernel component provided a weak
sink. This characteristic is probably associated with the long season and tropical origin of the Lfy germplasm.

Key words: Maize, grainfill, leafy, carbohydrate distribution, phenology, sink size

Andrews, C. J., Dwyer, L. M., Stewart, D. W., Dujas, J.-A. et Bonn, P. 2000. Répartition des hydrates de carbone durant la
phase de remplissage du grain chez les hybrides de maïs «feuillus» et chez les hybrides normaux. Can. J. Plant Sci. 80: 87–95.
Le génotype feuillu (Lfy) du maïs, qui porte des feuilles additionnelles au-dessus de l’épi par comparaison aux génotypes ordi-
naires (non feuillus), a produit à certains endroits un effet d’accroissement sur les rendements. Nous avons comparé la répartition
des hydrates de carbone chez trois hybrides Lfy (dont deux de type «staygreen» c.-à-d. conservant plus longtemps leur surface foli-
aire verte) et chez un témoin normal, Pioneer 3790, dans une région de saison courte (2800 UTC maïs) convenant aux hybrides de
précocité relative de 85 jours. Les sucres totaux et l’amidon étaient mesurés le 1er août 1990 à tous les étages du couvert cultural,
feuilles et tiges. Cette répartition était ensuite utilisée pour le calcul de la teneur en hydrates de carbone totaux dans le couvert
végétal dans deux années subséquentes à partir de prélèvements de feuilles et de tiges effectués à seulement trois ou quatre étages
du couvert à partir et au-dessus de celui de l’épi. Les composantes de l’épi, spathes, rafle et grain, étaient également analysées à
trois dates de prélèvement durant la phase de grossissement du grain. Des différences importantes étaient observées d’une année
à l’autre, mais d’une façon générale il y avait approximativement deux fois autant d’hydrates de carbone dans la partie du couvert
située au niveau et au-dessus de l’épi chez les lignées Lfy que chez le témoin ordinaire. En outre, les spathes et la rafle chez ces
lignées contenaient plus d’hydrates de carbone, mais en revanche en dessous du niveau de l’épi les concentrations étaient nette-
ment plus basses. Le taux de grossissement du grain était dans l’ensemble plus rapide chez les hybrides Lfy, mais chez l’hybride
Lfy A, le type à vieillissement précoce, il était plus lent durant l’année de culture plus fraîche (1992). Malgré la plus forte con-
centration d’hydrates de carbone observée chez les hybrides Lfy par rapport au témoin, leurs rendements grainiers n’en n’étaient
que légèrement supérieurs, ce qui laisse à penser que le grain serait un puits d’accumulation peu important, caractère probable-
ment associé à la longue saison de végétation et l’origine tropicale du type Lfy.

Mots clés: Hydrates de carbone, couvert végétal, maïs, génotype feuillu (Lfy)

The efficiency of light interception by the plant canopy is
one of the most critical factors in determining the yield of
maize. Leaf number, area, angle and orientation contribut-
ing to an avoidance of self-shading all may influence light
interception and thereby yield (Whigham and Woolley
87
1974; Tollenaar and Dwyer 1997). Leaf number is a major
component of leaf area, but amongst high-yielding hybrids
of the same maturity under the same conditions of tempera-
ture and photoperiod, variation in total leaf number is some-
what limited (Chase and Nanda 1967; Tollenaar and Hunter
88 CANADIAN JOURNAL OF PLANT SCIENCE

Table 1. Hybrids, leaf levels, and carbohydrate sampling dates in 1990, 1991 and 1992
Hybrids
Code Staygreen Pedigree Leaf Level
Lfy hybrid A No 2E5L × GL619A2 –5 to +10z
Lfy hybrid B Yes 1207A × GL619A2 –5 to +10
Lfy hybrid Cy Yes GL619A2 × 27 –5 to +10
Check Yes Pioneer 3790 –6 to +5

Sampling dates
1990 1991 1992
Date GDDx Date GDD Date GDD
S1 1 Aug 716 7 Aug 957 13 Aug 592
S2 – – 28 Aug 1204 2 Sept 743
S3 – – 17 Sept 1371 28 Sept 862
zLeaf levels are number of green leaves at pollination relative to ear leaf equal to zero.
yLfy C only studied in 1991 and 1992.
xGrowing degree days.

Table 2. Percentage distribution of biomass determined from 1 August 1990 sampling. Values are the means of six replications and two population
densities, as there were no interactions with density (data not shown)
Stem Leaves
Hybrid Portion Biomass (g) % Biomass (g) %
Lfy hybrid A Above ear 25.7 44.1 9.8 66.2
At ear 3.7 6.3 1.2 8.1
Below ear 28.9 49.6 3.8 25.7

Lfy hybrid B Above ear 42.3 56.5 14.0 74.2

At ear 4.3 5.8 1.3 6.7
Below ear 28.2 37.7 3.6 19.1

Check Above ear 10.3 21.4 5.6 45.9

At ear 2.5 5.2 1.3 10.8
Below ear 35.2 73.4 5.3 43.3

1983). An exception to this is found in Leafy (Lfy) maize
(“extra leaf maize”) described by Shaver (1983), which may
have six to eight extra leaves above the ear. A potential for
higher yields has been assumed for Lfy maize based on pho-
tosynthate from the additional leaves moving downwards to
the ear, and the higher average metabolic rate of the younger
leaves above the ear (Shaver 1983; Stewart et al. 1997).
In shorter seasons (85 d relative maturity), Lfy genotypes
did not provide the yield increases that were expected
(Dwyer et al. 1995), despite strong evidence (Hunter 1980;
Corke and Kannenberg 1989; Dwyer et al. 1994) that maxi-
mum leaf area, vegetative phase duration and duration of
leaf area are correlated with grain yield in these environ-
ments. There may be two reasons, proposed by Stewart et al.
(1997), for the failure of the Lfy plant type to have large
increases in grain yield. One is shading of the ear leaf region
known to contribute substantially to grain growth by the
extra leaves. In addition, the additional path length from the
extra source leaves to the grain may present a barrier to
sugar translocation within the stem, or at the stage of
unloading of the phloem to the grain.
A study was made of the performance of Lfy genotypes at
Ottawa, Canada, to gain information to better exploit the
potential for yield increases. Carbohydrate concentrations in
the canopy were found to be very different in the Lfy geno-
types than in a normal check hybrid, and expressions were
developed to relate late season decreases in canopy carbo-
hydrate concentrations to yield rankings (Dwyer et al.
1995). Input of such data and photosynthetic rates into a
model described growth of both Lfy and check hybrids, and
showed higher leaf export rate coefficients in the Lfy geno-
types (Stewart et al. 1997). The current report describes the
changing distribution of the total content of sugar and starch
in the canopy, the cob and husks and kernels during grain-
fill, and the relation of carbohydrate distribution to final
yield.
MATERIALS AND METHODS
Two experimental Lfy hybrids (A and B) and a normal
check hybrid (Pioneer 3790) were grown in 1990, and the
same hybrids and a third experimental Lfy hybrid (C) were
grown in 1991 and 1992. Hybrids and their leaf numbers are
listed in Table 1. Lfy hybrids B, C and Pioneer 3790 were
categorized as stay green with a longer duration for green
leaf area than the earlier senescing hybrid A. Plants were
grown on a sandy loam (Typic Haplorthod) in 1990, and on
a clay loam (Typic Eutrochrept) in 1991 and 1992 at
Ottawa, Ontario (45°23′N, 75°43′W). Planting density was
 ANDREWS ET AL. — GRAINFILL IN LEAFY AND NORMAL MAIZE
3.25 and 6.5 plants m–2 in 1990, and 4.0, 6.0 and 8.0 plants
m–2 in 1991 and 1992.). Hybrid by density combinations
were fully randomised within each block, and there were
three replications. Individual plots were of five row width in
1990 and 12 row width in 1991 and 1992.
Sampling Procedures
An intensive sampling of leaf and stem segments was taken
at each leaf level at five times throughout the day at anthe-
sis, 1 August 1990 and once early the next morning, to
establish biomass distribution and to determine single seg-
ment distribution of carbohydrate. In the next 2 yr, 1991 and
1992 plants were sampled at three dates, S1 to S3, during
grain filling. A sample consisted of two typical plants taken
from the outer six rows of the plot. On these dates, leaf and
stem segments were sampled late in the afternoon at four
levels in Lfy hybrids and three levels in the check (Table 1).
Cobs were sampled, husks and shanks were chopped, and
subsamples of leaf, stem, cobs and husk with shank were
accumulated over dry ice in the field and thereafter stored at
–20°C prior to analysis. The remaining tissues, with leaf and
stem segments bulked from all leaf levels, together with the
separated cob and husk with shank components were dried
at 100°C for biomass determination. Weight of the subsam-
ples taken for carbohydrate analysis was incorporated later.
Grain yield was determined from harvest of the two cen-
tre 7-m-long rows in October 1991 and November 1992 and
expressed at 155 g kg–1 moisture.
Carbohydrate Analysis
Samples were prepared and analysis carried out as described
by Dwyer et al. (1995). Total soluble sugar analysis was
based on the method of Gaines (1973), and starch on the
procedures of Reed and Singletary (1989) using a digestion
89
Fig. 1. Carbohydrate content of each leaf and
the associated stem segment for individual
leaf levels on 1 August 1990 for two Lfy and
one normal check hybrids. Plant density 6.5
plants m–2. (LSD0.05 between hybrids at com-
mon levels, 0.23.)
of starch to sugar by amyloglucosidase. Sugars in both sets
of samples were estimated by the phenol-sulphuric proce-
dure of Dubois (1965) using glucose as a standard. The
resulting sample concentration (mg g–1 dry wt) was multi-
plied by the weight of the corresponding plant part to give
carbohydrate content.
An estimate was made of carbohydrate content of intact
aboveground plants in 1991 and 1992 from the samplings
made from three or four leaf levels. The distribution of bio-
mass among all leaf levels, determined from the intensive
sampling of August 1990, was used to calculate percentage
biomass distributed at the ear node, below the ear and above
the ear (Table 2). Because leaf distribution of the three
Leafy hybrids is similar, a mean of the percentage biomass
distribution of Leafy A and B was used to calculate esti-
mates for Leafy C. These percentages were applied to total
stem and leaf biomass in 1991 and 1992 to determine bio-
mass at the ear node, below the ear and above the ear. Mean
carbohydrate concentration of these plant parts were then
used to calculate carbohydrate content.
Statistical Analysis
Analyses of variance were made each year at individual
sampling times on the sugar and starch contents of each
plant component, and the final kernel weight and grain
yield. Least significant differences (LSD 0.05) were calcu-
lated for significant effects. Correlations were calculated
between final grain yield and the carbohydrate contents of a
number of the plant components (n = 12, 4 hybrids, 3 densi-
ties).
RESULTS
Averages over the day of intensive leaf level sampling at
anthesis indicated that carbohydrate distribution was very
90 CANADIAN JOURNAL OF PLANT SCIENCE

Table 3. Contents of sugar and starch in stems and leaves below the ear and at the ear in three Leafy and one Check hybrid at three sampling
times in 1991 and in 1992
Below the ear At the ear
Stem Leaf Stem Leaf
Sugar Starch Sugar Starch Sugar Starch Sugar Starch
1991
S1 LfyA 14.69 1.10 0.94 0.19 2.00 0.14 0.32 0.07
LfyB 11.33 0.70 0.76 0.15 1.78 0.12 0.28 0.06
LfyC 13.09 0.83 0.94 0.18 2.00 0.11 0.31 0.07
Chk 18.31 1.07 1.81 0.35 1.51 0.06 0.22 0.12
LSD0.05 2.60 0.30 0.16 0.06 0.32 0.03 0.05 0.02

S2 LfyA 9.09 0.91 0.80 0.12 1.34 0.11 0.29 0.03

LfyB 8.29 0.55 0.70 0.09 1.31 0.08 0.27 0.03
LfyC 10.04 0.72 0.87 0.08 1.56 0.09 0.32 0.04
Chk 14.65 0.95 1.59 0.21 1.12 0.07 0.40 0.08
LSD0.05 3.06 0.33 0.17 0.05 NS 0.03 0.05 0.02

S3 LfyA 7.38 0.75 0.29 0.19 1.68 0.08 0.14 0.06

LfyB 8.38 0.49 0.31 0.14 1.47 0.08 0.14 0.05
LfyC 11.12 0.80 0.48 0.16 1.53 0.09 0.23 0.05
Chk 18.56 1.18 1.10 0.31 1.18 0.07 0.45 0.09
LSD0.05 7.51 0.34 0.41 0.50 NS NS 0.12 0.02

1992
S1 LfyA 22.29 2.24 1.05 0.18 3.39 0.26 0.38 0.08
LfyB 16.29 1.43 0.50 0.08 2.56 0.22 0.19 0.03
LfyC 17.25 1.25 0.97 0.16 2.52 0.22 0.36 0.07
Chk 24.10 2.40 1.64 0.29 1.73 0.14 0.38 0.10
LSD0.05 4.61 0.54 0.12 0.05 0.55 0.06 0.03 0.02

S2 LfyA 28.44 1.36 1.08 0.28 4.27 0.16 0.40 0.12

LfyB 16.47 0.72 0.57 0.14 2.72 0.15 0.23 0.07
LfyC 20.62 0.88 0.94 0.29 2.72 0.13 0.34 0.14
Chk 29.67 1.42 1.66 0.58 2.42 0.11 0.47 0.20
LSD0.05 4.25 0.27 0.16 0.05 0.48 0.03 0.07 0.03

S3 LfyA 23.23 1.30 1.29 0.16 3.11 0.15 0.48 0.08

LfyB 6.60 0.46 0.62 0.11 1.16 0.09 0.25 0.04
LfyC 20.56 0.96 1.31 0.20 2.63 0.14 0.47 0.06
Chk 25.69 1.34 2.07 0.31 2.04 0.09 0.58 0.08
LSD0.05 4.55 0.31 0.18 0.04 0.58 0.02 0.07 0.02
LSD0.05 = least significant difference at the 0.05 level of probability. NS = not significant at this level.

different in the Lfy genotypes compared to the check (Fig.
1). Major carbohydrate contents were found in the Lfy geno-
types at stem levels above those existing in the check.
Below the ear, carbohydrate contents of Lfy hybrid A and
the check were similar.
1991
At the ear level, there was consistently more sugar and
starch in the stems of the Lfy genotypes than the check at all
sampling times, but in the leaves at the ear, there was more
sugar and starch in the check (Table 3).
Below the ear, contents of sugar and starch in the leaf and
stem were higher in the check than the Lfy genotypes by a
factor of 1.5 to 2 (Table 3) while at and above the ear there
was approximately twice as much sugar and starch in the
Lfy genotypes as in the check (Fig. 2). The cob and the husk
component including the predominant mass of the shank
were together considered to be a transport pathway to the
kernels. Sugar and starch in this combined component
decreased from early to late sampling times and were lower
in the check than in the Lfy hybrids (Fig. 2).
With the decline through time of the carbohydrate of the
combined cob and husk component, that of the kernels
increased rapidly (Fig. 3). By S3, kernel carbohydrate of the
Lfy genotypes was greater than the check. Rate of kernel fill
(Table 4) was also greater in the Lfy hybrids.
At S3, there were significant interactions between hybrid
and plant density for both sugar and starch in the cob and
husk (Fig. 4). The reduction in carbohydrate of this compo-
nent due to density was much greater in the Lfy genotypes
than the check. Kernel yield was similarly reduced at the
higher density, but less so in the check than in Lfy (Fig. 5).
Kernel yield was greater in Lfy hybrid C than in the check,
which was reflected in final grain yields per hectare (Table
5). Grain yields of two of the Lfy genotypes were not dif-
ferent from the check.
 ANDREWS ET AL. — GRAINFILL IN LEAFY AND NORMAL MAIZE
1992
In most parts of the canopy, carbohydrate contents were
higher in the cooler year of 1992 than in 1991. At the ear
level there was more sugar and starch in the stem of Lfy
hybrid A than the other genotypes (Table 3). At the first two
sampling dates, the lowest stem sugar and starch were in the
check, but by S3 the lowest stem sugar, and the lowest leaf
sugar and starch were in Lfy hybrid B.
Below the ear, the Lfy genotypes contained less (>30%)
carbohydrate than the check in all cases, with the content of
Lfy hybrid B being substantially the lowest (Table 3). At
and above the ear, the canopy of the check contained less
91
Fig. 2. Contents of sugar and starch in the
upper canopy (stems and leaves at and above
the ear) and in the combined cob and husk with
shank component of three Lfy and one check
hybrids at three sampling times in 1991 and
1992. LSD0.05 between hybrids in stem sugar
in 1991: S1 = 2.8; S2 = 2.7; S3 = 4.9. In 1992:
S1 = 4.4; S2 = 3.9; S3 = 3.5. Other components
show significant (P < 0.01 to 0.001) differ-
ences between hybrids, with the exception of
1991 S2 leaf starch (P > 0.05) and 1991 S3 leaf
sugar, (P > 0.05). LSD0.05 between hybrids in
cob and husk sugar in 1991: S1 = 4.4; S2 = 2.8;
S3 = 2.5. In 1992: S1 = 3.1; S2 = 5.5; S3 = 1.8.
Cob and husk starch hybrid differences are sig-
nificant (P < 0.05 to 0.005) in both years with
the exception of 1991 S2 starch (P > 0.05).
sugar and starch than the Lfy genotypes, and by S3, hybrid
B contained the least sugar and starch of the Lfy hybrids
(Fig. 2). The carbohydrate of the combined cob and husk
component increased through the sampling times, in con-
trast to the response in 1991 (Fig. 2). Again, sugar and
starch contents of the check were lower than the Lfy
hybrids, with the exception of hybrid B at S3.
When the above components were compiled together
(Fig. 3), the total canopy (minus cob and husk) carbohydrate
was greater than in the previous year, but decreased marked-
ly by S3 in Lfy hybrid B. The accumulation of kernel car-
bohydrate was delayed in comparison with that in 1991, but
92 CANADIAN JOURNAL OF PLANT SCIENCE
Fig. 3. Total shoot carbohydrate of three Lfy and one check
hybrids at three sampling times. LSD0.05 of kernel component
between hybrids at S3 in 1991: 16.9; in 1992: 14.7. Kernels not
isolated at S1 in 1992. At S2, hybrid differences in kernel carbo-
hydrate not significant in both years. Other plant components sig-
nificant for hybrid differences in sugar and starch fractions as
described for Table 3 and Fig. 2.
was greater in hybrid B than the other genotypes. Rate of
kernel fill (Table 4) was greater in hybrid B and hybrid C
than in the check hybrid.
There were significant interactions between hybrids and
densities at S3 in cob and husk sugar and starch (Fig. 4).
This took a different pattern in 1992 than in the previous
year; these components decreased for all hybrids with
increasing density, but the relative decreases differed.
Kernel yield was reduced at the higher density and was
greatest in Lfy hybrid B (Fig. 5). Final grain yields were
substantially lower than in 1991, and there was less density
compensation than in 1991 (Table 5). The previous year’s
top yielding hybrid, Lfy hybrid C, together with hybrid A,
gave yields in 1992 significantly lower than the check, and
the grain yield of hybrid B was not significantly higher than
the check.
DISCUSSION
The 2 yr in which yield and carbohydrate data were collect-
ed and summarized in this report were remarkably different.
Table 4 . Rate of increase in carbohydrate in kernels of four hybrids
(grain fill rate) for two periods in 1991, one period in 1992 and with
means over all time intervals. Averages of three plant densities
Hybrid 1991 S1-S2 1991 S2-S3 1992 S2-S3z Mean
g plant –1 day –1
Lfy A 2.21 1.32 1.72 1.75
Lfy B 2.74 1.14 2.95 2.28
Lfy C 2.61 1.69 2.78 2.34
Check 2.11 0.53 2.35 1.66
LSD 0.05 (P = 0.33) (P = 0.16) 0.74 0.62
z1992 S1-S2 data are not presented because kernels were not present at
S1 sampling time.
The first year, 1991, was above average in heat unit accu-
mulation, but the second year was cooler than average with
greater rainfall. A delay in sampling by 1 or 2 wk in the
second year could not compensate for this difference in
growing degree days. Final harvest in 1992 was late, but
none of the hybrids under test achieved physiological matu-
rity.
The most marked difference between the 2 yr was the
overall greater size and greater accumulation of carbohy-
drate within the canopy in the cool growing season of 1992
than in 1991. The reasons for this are not clear. The mean
maximum temperature for July and August in 1991 was
27.3°C and for the same months in 1992 was 23.0°C; a dif-
ference of 4.3°C. This would be expected to lead to higher
photosythetic rates and greater photosynthate accumulation
in 1991, but the opposite actually occurred. This is in
aggreement with the work of Setter and Flannigan (1986)
and Tollenaar (1989) who showed that the optimum tem-
perature for dry weight accumulation in maize is between 19
and 23°C, despite the optimum for photosynthesis being
27°C. In 1991, field temperatures were frequently above this
optimum, with high potential evapotranspiration, both act-
ing to substantially reduce photosynthetic activity. There
were also observations of mid-summer water deficit stress
in 1991, including afternoon leaf wilting, particularly at
high population density.
Further, in 1991, there was an earlier and greater accu-
mulation of carbohydrate in the kernels than in 1992. This
rapid accumulation takes place at the same time as the
depletion of carbohydrate in the rest of the canopy. In con-
trast, in 1992 grainfill occurred later, but with the exception
of Lfy hybrid B, was not accompanied by a loss of canopy
carbohydrate, presumably because of a continuing high rate
of photosynthetic accumulation. The case of hybrid B will
be discussed later.
A consistent difference emerging from this study is that
the Lfy genotypes have about twofold more carbohydrate in
the upper canopy than the check. This occurs in the stem,
leaves and to a lesser extent in the cob and husk component.
This large source does not result in dramatically higher
grainfill rates or kernel weight. A counteracting factor is
undoubtedly that below the ear, the check hybrid accumu-
lates substantially more carbohydrate than the Lfy types.
The role of this pool of carbohydrate below the ear is
 ANDREWS ET AL. — GRAINFILL IN LEAFY AND NORMAL MAIZE
Fig. 5. Kernel harvest yield, per plant, at three planting densities,
4.0, 6.0 and 8.0 plants m–2. LSD0.05 between hybrid means, in
1991: 17.4, in 1992: 12.3.
93
Fig. 4. Sugar and starch contents of the
cob and husk component of three Lfy
and one check hybrids grown at three
planting densities 4.0, 6.0, and 8.0 plants
m–2 at the S3 sampling. Significant den-
sity × hybrid interaction. LSD0.05 in
1991: sugar = 4.4, starch = 0.7; in 1992:
sugar = 3.2, starch = 1.4.
uncertain and information on its significance depends on the
nature of the study. Conclusions range from no contribution
to grain growth (Fairey and Daynard 1978), to 30% in trop-
ical maize (Palmer et al. 1973) and 50% in temperate sweet
corn (Sawada et al. 1995). If remobilization or transport of
carbohydrate from the lower canopy to the grain is as sub-
stantial as these studies indicate, it is reasonable to suppose
that the greater pool in the lower canopy of the check would
contribute significantly to grainfill and counteract the defi-
ciency (in relation to the Lfy) in the amount of carbohydrate
available from the upper canopy. It is perhaps significant in
this regard that the difference between Lfy and check
hybrids in carbohydrate of the cob and husk component is
considerably smaller than the difference in the upper canopy
in both years. This change could be explained by additional
carbohydrate from the lower canopy in the check.
The Lfy hybrids generally have faster grainfill rates than
the check, with the exception of hybrid A, the early senes-
cent hybrid in 1992. In an attempt to extract a comparison
between the two very different years, days after pollination
at S3 in the cool year 1992 were taken and carbohydrate
contents at the equivalent time in 1991 interpolated (Fig. 6).
In each hybrid at this equivalent developmental stage, grain-
fill was further advanced in 1992 than in 1991, and was
particularly advanced in hybrid B. This indicates that there
is a complex relationship between growing degree days,
chronological days and development and grainfill in these
hybrids.
Of the Lfy genotypes, hybrid B shows a different distrib-
ution of carbohydrate during the grainfill period, particular-
ly noticeable in the cooler season of 1992. In that year, by
S3, carbohydrate content of the canopy at the ear level and
above, including the cob and husk, was lower than the other
94 CANADIAN JOURNAL OF PLANT SCIENCE

Table 5. Final grain yield of four hybrids at three planting densities in 1991 and 1992
1991 1992
Lfy A Lfy B Lfy C Check LfyA Lfy B Lfy C Check
t ha–1
Plants m–2
4 7.67 9.47 9.85 8.22 5.92 6.71 4.49 5.46
6 9.36 9.95 11.09 9.09 4.70 6.77 3.66 6.46
8 9.56 9.05 11.75 10.34 4.56 6.95 4.58 6.20

Mean 8.86 9.49 10.90 9.22 5.06 6.81 4.24 6.04

LSD0.05 between hybrid means in 1991 = 1.16; in 1992 = 0.85

B has greater transport capacity from source to sink, that is

Fig. 6. Carbohydrate content of plant components of three Lfy and
one check hybrid at equivalent days after pollination in 1991 and
1992. Data for 1992 fixed at S3; 1991 data interpolated.
Lfy genotypes. Kernel fill proceeded more rapidly than the
other genotypes, and hybrid B attained the highest yield.
Photosynthetic rates of hybrid B are similar to those of
hybrid A, and the leaf export coefficients are similar
between Lfy hybrids and higher than those of the check
(Stewart et al. 1997). The observations suggest that hybrid
particularly sensitive to cooler temperatures.
Carbohydrate contents of a number of plant parts were
correlated with final kernel yield in both years (Table 6). In
1991, final kernel yield was correlated with the canopy car-
bohydrates and with carbohydrates of the stem at the ear. No
associations below the ear were found, but there were high
correlations between kernel yield and the contents of all the
aboveground stem, and with the total aboveground plant.
These relationships are a reflection of the correlations
between biomass at silking and final yield described previ-
ously in short-season maize (Daynard and Kannenberg
1976; Dwyer et al. 1994).
The Lfy phenotype has several advantages over the nor-
mal maize plant. It has greater leaf area, which is developed
rapidly. In the hybrids investigated here, greater leaf area
does not delay the onset of grainfill, which has been an
obstacle to improvement of short-season corn (Cross 1975;
Hunter 1980; Corke and Kannenberg 1989). The Lfy lines
have higher photosynthetic rates and higher leaf export rates
(Stewart et al. 1997), thus minimizing carbohydrate storage
in the leaf, but overall there is a higher concentration of car-
bohydrates in the upper canopy (Dwyer et al. 1995). The
current data show that there is about twice as much carbo-
hydrate in the upper canopy in the Lfy than the normal
maize hybrid. Despite the fact that in these studies Lfy lines
did inconsistently outyield the check [Dwyer et al. (1995)
and Table 5], it is apparent that they did not maximize the
use of available carbohydrate.
The model described by Stewart et al. (1997) predicts the
enhanced movement of carbohydrate in Lfy from leaves to
stem, but does not extend to movement through the stem or
into the cobs and kernels. Content of carbohydrate in the
cobs and husks of the Lfy lines is greater than that of the
check. This supports the view that the kernels of the Lfy
hybrids constitute a weak sink, which cannot accommodate
all the available carbohydrate. In contrast, the yields of
short-season corn are frequently limited by an inadequate
supply of carbohydrate during grainfill (Tollenaar 1977;
Corke and Kannenberg 1989). A weak sink is a common
feature in tropical and extended season maize, and indica-
tions of a weak sink in the Lfy hybrids of this study may be
a reflection of the long-season background of the Lfy types
(Shaver 1983).
 ANDREWS ET AL. — GRAINFILL IN LEAFY AND NORMAL MAIZE 95

Table 6. Correlations between final kernel yield and carbohydrate contents in a number of aboveground plant components
Date Kernel Cob/husk/shank Canopy Stem Stem Stem Plant
Total Total At ear Below ear Total Total
CHO CHO CHO CHO CHO CHO CHO
1991 kernel yield S1 0.462 NS 0.867** 0.596* 0.773** 0.398 NS 0.815** 0.842**
S2 0.754** 0.742** 0.555 NS 0.695* 0.504 NS 0.738** 0.736**
S3 0.840** 0.792** 0.615* 0.655* 0.472 NS 0.727** 0.750**

1992 kernel yield S1 – 0.625* 0.140 NS 0.296 NS 0.319 NS 0.334 NS 0.331 NS

S2 0.648* 0.765** 0.438 NS 0.460 NS 0.459 NS 0.642* 0.633*
S3 0.660* 0.261 NS 0.095 NS -0.111 NS 0.055 NS 0.087 NS 0.095 NS
*,** Correlation significant at 0.05 level of probability and 0.01 level of probability, respectively (n = 12, 4 hybrids × 3 densities). NS, no significant cor-
relation.

Chase, S. S. and Nanda, D. K. 1967. Number of leaves and matu-
rity classification in Zea mays L. Crop Sci. 7: 431–432.
Corke, H. and Kannenberg, L. W. 1989. Selection for vegetative
phase and actual filling period duration in short season maize. Crop
Sci. 29: 607–612.
Cross, H. Z. 1975. Diallel analysis of duration and rate of grain-
filling of seven inbred lines of corn. Crop Sci. 15: 532–535
Daynard, J. B. and Kannenberg, L. W. 1976. Relationships
between length of the actual and effective grain filling periods and
the grain yield of corn. Can. J. Plant Sci. 56: 237–242.
Dubois, M., Gilles, K. A., Hamilton, J. K., Rebers, P. A. and
Smith, F. 1965. Colorimetric method for determination of sugars
and related substances. Anal. Chem. 28: 350–356.
Dwyer, L. M., Andrews, C. J., Stewart, D. W., Ma, B. L. and
Dugas, J.-A. 1995. Carbohydrate levels in field-grown Leafy and
normal maize genotypes. Crop Sci. 35: 1020–1027.
Dwyer, L. M., Ma, B. L., Evenson, L. and Hamilton, R. I. 1994.
Maize physiological traits related to grain yield and harvest mois-
ture in mid- to short-season environments. Crop Sci. 34: 985–992.
Fairey, N. A. and Daynard, T. B. 1978. Assimilate distribution
and utilization in maize. Can. J. Plant Sci. 58: 719–130.
Gaines, T. P. 1973. Automated determination of reducing sugars,
total sugars and starch in plant tissues from one weighed sample. J.
Assoc. Off. Anal. Chem. 56: 1419–1424.
Hunter, R. B. 1980. Increased leaf area (source) and yield of
maize in short season area. Crop Sci. 20: 571–574.
Palmer, A. F. E., Heichel, G. E. and Musgrave, R. B. 1973.
Patterns of translocation, respiratory loss and redistribution of 14C
in maize labelled after flowering. Crop Sci.13: 371–376.
Reed, A. J. and Singletary, G. W. 1989. Roles of carbohydrate
supply and phytohormones in maize kernel abortion. Plant Physiol.
91: 986–992.
Sawada, O., Ito, J. and Fujita, K. 1995. Characteristics of photo-
synthesis and translocation of 13C-labelled photosynthate in husk
leaves of sweet corn. Crop Sci. 35: 480–485.
Setter, T. L. and Flannigan, B. L. 1986. Sugar and starch redis-
tribution in maize in response to shade and ear temperature treat-
ment. Crop Sci. 26: 575–579.
Shaver, D. 1983. Genetics and breeding of maize with extra leaves
above the ear. Pages 161–180 in Proceedings of the 38th Annual
Corn and Sorghum Industry Res. Conf. Chicago IL. Am. Seed
Trade Assoc. Washington, DC.
Stewart, D. W., Dwyer, L. M., Andrews, C. J. and Dugas, J.-A.
1997. Modelling leaf carbohydrate production, storage and export
in Leafy and normal maize (Zea mays L). Crop Sci. 37:
1228–1236.
Tollenaar, M. 1977. Sink–source relationships during reproduc-
tive development in maize. A review. Maydica 22: 49–75.
Tollenaar, M. 1989. Response of dry matter accumulation in
maize to temperature. I. Dry matter partitioning. Crop Sci. 29:
1239–1246.
Tollenaar, M. and Dwyer, L. M. 1997. Physiology of maize. In
D. L. Smith and C. Hamel, eds. Crop physiology. Springer Verlag,
New York, NY.
Tollenaar, M. and Hunter, R. B. 1983. A photoperiod and tem-
perature sensitive period for leaf number of maize. Crop Sci. 23:
457–460.
Whigham D. K. and Woolley, D. G. 1974. Effect of leaf orienta-
tion, leaf area and plant densities on corn production. Agron. J. 66:
482–486.